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A New Capitosaur from the Middle Triassic of Spain and the Relationships Within the Capitosauria

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A New Capitosaur from the Middle Triassic of Spain and the Relationships Within the Capitosauria A new capitosaur from the Middle Triassic of Spain and the relationships within the Capitosauria JOSEP FORTUNY, ÀNGEL GALOBART, and CARLES DE SANTISTEBAN Fortuny, J., Galobart, À., and De Santisteban, C. 2011. A new capitosaur from the Middle Triassic of Spain and the rela− tionships within the Capitosauria. Acta Palaeontologica Polonica 56 (3): 553–566. Capitosaurs were the largest and homogeneous group of Triassic temnospondyl amphibians with cosmopolitan distribu− tion. However, their interrelationships are debated. The first capitosaur cranial remains found in the Iberian Peninsula were assigned to Parotosuchus; herein, a re−description of this material, together with information on other remains re− covered from the same site, enables us to classify them as a new genus: Calmasuchus acri gen. et sp. nov. (Amphibia: Temnospondyli) from the early−to−middle Anisian (early Middle Triassic). This capitosaur had a combination of plesiomorphic and non−plesiomorphic characters, such as posterolaterally directed tabular horns, paired anterior palatal vacuities, and unique morphology of the lower jaw. By cladistic analysis, we propose a new phylogeny for the monophyletic capitosaurs. In the analysis, Capitosauria is supported by seven synapomorphies. Wetlugasaurus is the most basal member of the clade. The score of the Russian taxon Vladlenosaurus alexeyevi resulted in a clade including Odenwaldia and the latter taxa. The Madagascarian Edingerella is the sister taxon of Watsonisuchus. Finally, Calma− suchus acri, the new taxon described here, appears as a more derived form than Parotosuchus. The new genus is the sister taxon of the Cyclotosaurus–Tatrasuchus and Eryosuchus–Mastodonsaurus clades. Key words: Temnospondyli, Capitosauria, Mastodonsauroidea, phylogeny, computed tomographic scanning, Anisian, Triassic, Spain. Josep Fortuny [[email protected]] and Ángel Galobart [[email protected]], Institut Català de Paleontologia, Universitat Autònoma de Barcelona, Edifici ICP, Campus de Bellaterra s/n, 08193 Cerdanyola del Vallès, Barcelona, Spain; Carles de Santisteban [[email protected]], Departament de Geologia, Universitat de València, C/o Dr. Moliner 50, 46100 Burjassot, València, Spain. Received 12 March 2010, accepted 20 December 2010, available online 29 December 2010. Introduction The first revision of the group (Welles and Cosgriff 1965) focused on the evolution of the otic notch (open in primitive Capitosaurs constituted one of the most abundant groups of forms and closed in derived forms) in an attempt to establish Triassic temnospondyl amphibians. Despite their cosmo− the taxonomy of the “Capitosauridae”. The revision by Ochev politan distribution, fossil records in the Iberian Peninsula (1966) was markedly different: a large number of genera and are scarce, with only three reported sites: Rocha da Pena, species were recognised by using the shape of the tabular horn Portugal (Witzmann and Gassner 2008); and La Mora and as a criterion for determining taxonomic relationships. The Riera de Sant Jaume, Spain (Gaete et al. 1993; JF unpub− most recent revisions (Schoch 2000; Damiani 2001; Schoch lished data). 2008; Maganuco et al. 2009) demonstrate that the evolution of The wide distribution of temnospondyls, especially capito− the group involves more cranial characters than those regarded saurs, has been used in biostratigraphy (Cosgriff and Defauw previously. 1987) to correlate Triassic tetrapod faunas (Lucas 1998). On the other hand, recent papers include new descriptions These faunas are well known in the Eastern European plat− of old and new specimens recovered from the Central Euro− form (e.g., Ochev and Shishkin 1989; Shishkin and Ochev pean Basin, providing crucial information to understand the 1994) and Beafourt Group of South Africa (Karoo Basin) evolution of the clade; especially, capitosaur remains have (e.g., Hancox et al. 1995; Shishkin et al. 1995; Damiani 2004). been described in detail (Maryańska and Shishkin 1996; However, the phylogenetic relationships of capitosaurs have Schoch 1997, 1999; Sulej and Majer 2005; Shishkin and remained controversial since the first discoveries, even with Sulej 2009). The first capitosaur cranial remains recovered in the more recent studies (e.g., Watson 1962; Welles and Cros− Spain were assigned to Parotosuchus (Gaete et al. 1993, griff 1965; Ochev 1966; Schoch 2000; Schoch and Milner 1994, 1996). Here, we present a revised description of the 2000; Damiani 2001; Steyer 2003; Schoch 2008; Maganuco et original and new materials, enabling us to establish a new ge− al. 2009). nus—Calmasuchus acri gen. et sp. nov. (Amphibia: Temno− Acta Palaeontol. Pol. 56 (3): 553–566, 2011 http://dx.doi.org/10.4202/app.2010.0025 554 ACTA PALAEONTOLOGICA POLONICA 56 (3), 2011 Facies: A D C S4 A E D B D A D El Muntanyá C S3 Collformic cross stratifications SERRA DE PICAMENA C stream mark A E D C S2 Cenozoic Aiguafreda la Calma D B Buntsandstein Facies D E Lower Muschelkalk Facies B D S1 Middle Muschelkalk Facies A D Upper Muschelkalk Facies Tagamanent N 1m Paleozoic Paleozoic 1:50.000 0 0.75 1.50 km Fig. 1. Geographic and geologic location of the capitosaur Calmasuchus acri gen. et sp. nov. from the Anisian at the La Mora site, Spain, with a stratigraphic cross−section containing the vertebrate locality (arrow). See text for facies interpretation. spondyli)—and providing new data to resolve the evolution continental deposits (Calvet and Marzo 1994). These materi− of the group. In this regard, new data acquired by computed als are derived from the erosion of the Iberian Massif and tomography, including complete skull reconstruction of this they show thickness increment from northeast (145 m) to taxon, have been added to the previous information obtained southwest (310 m). by mechanical preparation of the matrix. Cladistic analysis The sedimentary deposits of this sub−basin are mainly including the new taxa performed by using an updated ver− composed of sandstone, mudstone, and red clay (Areniscas y sion of the previous matrix of Damiani (2001) for the group Lutitas del Figaró unit); these materials are interpreted as flu− has enabled us to propose a new phylogeny. vial deposits and distributed from northwest−northeast to southeast−southwest (Calvet and Marzo 1994). Institutional abbreviations.—IPS, Institut Català de Paleonto− At the La Mora site, the Buntsandstein facies deposits be− logia, Sabadell, Spain; MB, Museum für Naturkunde, Berlin, long to the lower part of the Areniscas y Lutitas del Figaró Germany; MNHN, Muséum National d'Histoire Naturelle, unit. This unit is unconformable on Paleozoic metamorphic Paris, France; PIN, Paleontological Institute of the Russian rocks. The section is 25 m thick and is composed of intercala− Academy of Sciences, Moscow, Russia. tions of red clay, mudstone, and sandstone in four sedimen− tary sequences with five different facies bound by caliche− type palaeosols (Fig. 1). The materials are interpreted as in− Geological setting filling of a channel body with an erosional base and a topping plane. Facies A represents the first deposits that formed the The Triassic outcrops of the La Mora site (Catalonia, Spain, base of the level and deposits B are enclosed by the margin of northeast of the Iberian Peninsula) belong to the Montseny− the channel. Both are interpreted as point bars. In relation to Llobregat domain (Calvet and Marzo 1994) of the Cata− these point bars, the sandstone from facies C fills the central lonian basin. This palaeogeographic unit has been identified part of the channel and its equivalent to the lower part of the as one of the eastern−most sub−basins of the Iberian plate. deposits of the point bar. Facies C yields the vertebrate local− The Buntsandstein facies in this area (Fig. 1) is formed by ity. The clay and silt from facies D constitute the final infill− FORTUNY ET AL.—NEW CAPITOSAUR FROM SPAIN AND CAPITOSAUR RELATIONSHIPS 555 50 mm Fig. 2. Computed tomographic reconstruction of the capitosaur Calmasuchus acri gen. et sp. nov. from the Anisian at the La Mora site, Spain, in dorsal (A), ventral (B), lateral (C), and occipital (D) views. ing of the abandoned fluvial channel and extend beyond the of this facies and the ammonite Paraceratites fauna is lo− boundary of the channel as flood−plain deposits. In facies E, cated in the middle part of the lower Muschelkalk facies (see the sediments show a concentration of calcium carbonate Márquez−Aliaga et al. 2000; Dinarès−Turell et al. 2005 for nodules. further details). All these data restrict the La Mora site to an The age of the Areniscas y Lutitas del Figaró unit remains Aegean−to−middle Pelsonian age (early−to−middle Anisian). uncertain. The first palynological work in this Buntsandstein facies (Solé de Porta et al. 1987) dated the unit as Anisian. A recent palaeomagnetostratigraphic study excludes the possi− Materials and methods bility of an Olenekian stage (Dinarès−Turell et al. 2005) and biostratigraphic analysis of the overlying lower Muschelkalk The La Mora site was discovered in 1989 by local hikers facies indicates an age of middle−to−upper Pelsonian to upper Emili Ramon and Pere Font. A year later, palaeontological Illyrian (middle−to−upper Anisian), on the basis of the pres− fieldwork was undertaken by the Institut de Paleontologia de ence of the conodonts Paragondolella bulgarica, P. han− Sabadell (Spain), extracting hundreds of cranial and post− bulogi, P. bifurcata, Neogondolella constricta, N. cornuta, cranial bones. Most of these bones were identified as
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